3/14 - Module 3, From Tree Shrew to Ape

Cenozoic Era:
• Overall cooling (with fluctuations)
• Changing temperatures caused by plate tectonics & ocean currents: Cenozoic era starts here

Paleocene Era:
Angiosperms had just had an adaptive radiation
– Flowering plants
• Nectar attracts animals that inadvertently carry pollen to other plants
– Evolved new, large fruit types
• Animals eat them and inadvertently spread seeds
– Became dominant tree species 100-60 Ma (replacing gymnosperms)

Pre-Angiosperm Forests
• Dominated by non-flowering trees (Gymnosperms)
• Pollen spread by wind
• Hot
• Open
• Not particularly complex
• Dominant animals: reptiles

Angiosperm Forest
• Lush, many layers
• Closed
• Cooler
• Many niches
• Dominant animals: mammals & birds
Example: fruit, orange, any fruit you eat

Angiosperm hypothesis of primate origins

• The availability of abundant fruits and flowers in the terminal branches of tropical forest trees
provided a windfall of foods (fruit AND insects) that were eaten by the earliest primate ancestors.

Plesiadapiforms
65-54 mya
Diverse, successful group, found in North America and Europe during Paleocene.
• 35 genera, 75 species
• 20g – 3 kg
• insectivorous & frugivorous
• Earliest primates?
• some similar traits to later primates
• molar shape, grasping hands
• one species had a nail on big toe
• but, had many non-primate traits, like … Toe bones
Eocene Era is the primarily era of where primate evolution occurred or began.
Paleocene-Eocene Thermal Maximum (PETM)
• Global temperatures rose dramatically at the beginning of the Eocene
• Many terrestrial mammals went extinct, but numerous modern
mammalian orders emerged

Environmental factors in the Eocene

• Whole world became warmer
Northern latitudes had tropical climates
• Tropical climates caused rain forests in North America, up to present-day Alaska
- primate movement across the world

At the beginning of the Eocene, Big Horn Basin was a swamp

Characteristics of Eocene Primates

• Postorbital bar
• Shorter snouts than other mammals
• Forward-facing eyes
• Nails instead of claws
• Opposable first digits

>200 species of Eocene primates

Two major groups (north america):

• Adapiformes
• Resembled lemurs
– Generalized arboreal quadrupeds with some leaping
– Folivorous
– Diurnal

• Omomyiformes
• Resembled tarsiers
– Insectivorous
– Nocturnal
– Leapers
And in Asia. . .
• Eosimiidae
• tiny primates
• Possibly ancestors of Haplorrhines
• 2 species ~10 grams

Climate Change from Eocene to Oligocene

• Surface sea water much cooler
• Antarctic ice cap developed
• Northernmost tropical forests disappeared

Emergence of Strepsirrhini and Haplorrhini

Haplorrhine Primates in the Fayum, Egypt (A place rich in Oligocene fossils)

• Earliest unambiguous Haplorrhines 36-33 Ma
• Reduction of snout compared to strepsirrhines
• Postorbital closure
• Mandibular fusion

Aegyptopithecus

Characteristics
– Generalized molars
– Racoon-sized
– Climber

• One of the earliest catarrhines*

• Came before split between Old world monkeys and Apes *group containing OWMs and Apes

How did they get there?

The scenarios:
1. Evolved right there in S. America: Probably not. No pre- Oligocene primate-like fossils in S.
America

2. Rafted from North America: Probably not.

Ocean currents:
-North AmericaEurope
-Africa South America

3. Rafted from Africa – a Fayum ancestor: Likely! Caviomorph rodents appear in S. America at
about the same time (They have African roots too)

Pirate Monkeys: new kids show idea?

All living New World Monkeys are descendants of this dispersal event!
Climate change again!
• Warming at end of Oligocene going into the Miocene
• Northern tropical forests returned

Proconsul species
• Genus found in about 6 localities
• Ranges from 20 to 17 mya
• Several species known
• Tail or no tail?
• Y-5 molar pattern (ape-like)
• Largish brain (ape-like)
• Flexible shoulder (ape-like)

Dryopithecus
• West-central Europe
• Short vertebral column
• Broad chest, long arms
• Powerful hands
Oreopithecus
East Africa and Southern Europe
• Late Miocene (8mya)
• Found in Tuscany
• Long arms/short legs
• Short trunk
• Mobile joints

Possibly bipedal

Inner ear bony labyrinth consistent with bipedalism and foot shows unique adaptation for
stability on two feet: a very widely displaced big toe

Sivapithecus
• Found in South Asia
• First appear around 13mya
• Disappear around ~5-8mya
• 3 species ascribed to genus Sivapithecus
• Generalized quadrupedal ape

Later discoveries suggest that Sivapithecus was related to the Orangutan

What About Africa?

• Gap in the fossil record
• 15 – 8 Ma

Darwin argued for an African origin of humans

● The origin of the human lineage is likely found in Africa, not Asia or elsewhere
● Molecular phylogenies indicate that the divergence of human ancestors from African
apes occurred 4-8 million years ago
● Chimpanzees & bonobos are the most recent living relative of humans

3/16/23

Mid/late Miocene
(15-8 ma):
• Ape diversity decreasing everywhere
• Few suitable fossil sites in Africa in general
• Very few (poorly preserved) African ape fossils
How to Make a Fossil
Phase 1: Death
– How an animal dies determines much of its future as a possible fossil

Phase 2: Burial

Phase 3: Replace calcium with minerals

Phase 4: Exposure
– Erosion
– Usually combined with tectonic activity

How do we know that “Lucy” lived around 3.2 mya?

Geologic dating methods

-Over time, radioactive isotopes (parent) change into stable isotopes (daughter)
-Geochronologists know various rates of isotopic decay
-Change is constant and can be expressed as a half life: the time it takes to reduce apparent
atoms in the material by 50%

Potassium-argon dating
● Feldspar is a mineral containing Potassium (K40 )
● K40 decays to Argon (A40 )
● Volcanic ash contains feldspar
● When a volcano erupts, K40 resets to 100%:
● Over time, K40 changes to Ar 40 at a known rate

Volcanic layer = Tuff

Paleomagnetism
-The earth’s magnetic polarity switches every few hundred thousand years. Iron particles in
rocks track this switch.

Faunal correlation (don’t need to know much about this)

-Some lineages of animals leave behind a good fossil record scattered over a wide area
-Get one absolute date for pigs in locality X.
-Pigs and hominid fossils are found in the same layer in locality Y
-Can estimate age of hominid fossil

What is a hominin?

• Family Hominidae: humans and all great

apes. a.k.a. Hominids.

• Subtribe Hominina: bipedal hominids

– These are called Hominins.

What are the skeletal traits that indicate bipedality?

-Position of Foramen Magnum (the hole through which the backbone attaches to the bottom of
the skull)

-Barrel-shaped rib cage (Chimp skeleton has cone shaped torso)

(Humans have more parallel sides→ helps us move our arms in easy fashion way)

-Verbal Column (gravity is know acting parallel to the spine rather than perpendicular)

-Curves of Vertebral Column

Humans→ Lordosis curve at end of spine

Lordosis affects center of gravity

Apes (in bipedal posture):

• Center of gravity in front of hips
• Constantly resisting tendency to fall forward

Humans:
• Center of gravity over hips
• Less energy resisting tendency to fall forward

Pelvis Shape

Iliac Blades
• Iliac blades are shorter and wider in humans (think of a bucket)
• “Basin-shaped” pelvis supports abdominal organs

Limb Proportions
Humans:
• Legs longer than arms
• Efficient bipedal stride

Apes:
• Arms longer than legs
• More efficient in trees

Intermembral Index (assign a number to how long an individual's legs are to their arms)
IMI = 70
IMI = 106
Forelimb length/Hindlimb length X 100

Valgus Knee

Arch in Foot
-Shock absorber
(all bones connected to ligaments that stretch)
Most humans have somewhat of an arch
Chimps are more flat foot

Convergent Big Toe

-Humans have a big toe that converged on the same plane as the other toes (parallel)
-Chimps have a big toe that is set out at an angle, this allows them to grasp with their feet

Straight toes bones

Humans- straight
Chimps- curved, less efficient for bipedal walking

Signatures of bipedalism:
-Position of foramen magnum
• Curvature of spine and sacrum
• Barrel-shaped rib cage
• Broad and shortened pelvis
• Valgus angle at knee joint
• Convergent big toe
• Arch in foot
• Intermembral index indicating long legs
• Straight toe bones

Are there advantages to being bipedal?

Why did bipedalism evolve?

1. More than just adapting to a terrestrial niche

2. (Provisioning model)
● Males would gather food and bring it back to females – hence, their hands needed to be
free to carry these items
-But, this would mean that pair-bonding evolved early
-Sexual dimorphism suggests otherwise...
3. Using tools model
● Hands needed to be free to make and use tools to hunt
-but don’t find evidence of stone tools until long after bipedalism
4. Thermoregulatory model
● Reduced heat gain
● Increases heat loss
5. Climbing Mechanical Model
● Retention of suspensory morphology
● Perhaps arboreal habitat became patchy and arboreal ancestors had to travel on the
ground and had to walk bipedally because of their morphology

But what about the fact that close hominin relatives knuckle walk?
● Comparative anatomy of living African apes and fossil hominins suggests that humans
evolved from a knuckle walking ancestor

WHAT CAUSED NATURAL SELECTION TO ACT ON THIS TYPE OF LOCOMOTION?

-we don’t know

Really general characteristics of early hominins

• Bipedal
• Slightly increased cranial capacity (brain size)
• Trend toward non-honing premolars

What is a honing premolar?

A pre-molar that sharpens upper canine

(Monkeys and apes)

Hominins: Trend toward non-honing premolar & smaller canines

(reduction of size)

Possible Early Hominins

• Range in time from 7 – 4 Ma
• Retain some primitive characteristics
• BUT
– Have some hominin features
• Four species

North Central Africa, Chad (the oldest specimen)

• Now desert
• Previously lakes
• 6 – 7 Ma
• A fossil skull

Sahelanthropus tchadensis
• Meaning of name
– sahel = African region
– Anthropus = human
– Tchadensis = from Chad
• Canine-premolar complex reduced (becoming non-honing)
• Biped (?)
– Foramen magnum

East African Rift Valley (huge component)

• Location
– Malawi through Eritrea and into the Red Sea

Orrorin tugenensis
• Meaning
– Orrorin = original man
– tugenensis = from Tugen Hills
• Biped
– femur shape
– But, curved phalanges
• Reduced canine
• Less-honing premolar
• 6.2-5.8 ma

The Middle Awash:

Ardipithecus kadabba
• Name
– ardi = ground
– pithecus = ape
– kadabba =original
-5.55 -5.2 Ma
Reduced canine, 3rd premolar has small honing facet

Ardipithecus ramidus
• Name
– ardi = ground
– pithecus = ape
– ramidu = root
• Dates
– 4.8 – 4.4 Ma

Ardipithecus ramidus
• Characteristics:
– Premolar less honing than in Ar. kadabba
– Bipedal, but has a divergent big toe
– Really long fingers

Australopithecus and Paranthropus

Australopithecus - early hominin that originated in Africa during Pliocene and Early Pleistocene.
- Skull/brain slightly bigger
- Strong nasal prognathism (“sticking-out” below nostril), no true function to this change
- Robust mandible (stronger,thicker,higher jaw bone), where we start to see dietary shift
- Canine Diastema - A space b/t upper incisor and upper canine to fit lower canine.

Australopithecus Anamensis: oldest one

- Name
- Australo: southern
- Pithecus: ape
- Anam: lake in Turkana language
- AGe: 4.2-3,8 Ma
- Oldest Au. Species
- Meave Leaky

Australopithecus Afarensis: little younger

- Name
- Australo: southern
- Pithecus: ape
- Afar: region and people of Ethiopia
 - Age: 3.8-2.95
- Don Johanson (ASU) discovered a knee joint at Hadar, Ethiopia in 1973. Inward slanted
knee joint is indicative of knees under the hips.
- Don Johanson also discovered Lucy in 1974 at Hadar.
- Lucy: Female AU Afarensis, she is small!

Kenyanthropus Platyops
- Name
- Kenya: the country
- Anthropus: human being
- Platyops: flat face
- Age: 3.5-3.2 Ma

Australopithecus Africanus (where southern ape comes from, first found of this genus)
- Name
- Australo: southern
- Pithecus: ape
- Africanus: Africa
- Age: 2.8-2.2 Ma?
- Recovered from cave sites in SOuth Africa
- **important cave finding**
- Suggests that bipedality came before increase in brain sides

Australopithecus Garhi
- Name
- Australo: southern
- Pithecus: ape
- Garhi: surprise
- Age: 2.5 Ma
- Big teeth, small brain
- Berhane Afshaw

Australopithecus Sediba
- Name
- Sediba: natural spring
- Age: 1.9 Ma
- Young son found a skull while running around in a cave.
- Small teeth

Ecology of Australopithecus
Diet
- In comparison to Miocene Apes, Australopithecus has large, flat molars with thick
enamel
 - Good for crushing hard, brittle items.
- 2 Carbon Isotopes (C4: Grasses, C3: Trees and Shrubs) in the area. This can be
detected in tooth enamel of an organism.

Phytoliths from the dental calculus

- (plaque) of Australopithecus sediba.
- • Microscopic minerals in plant cells
- • Each plant species has unique patterning
- • Re-construct diet

Stone-tool-inflicted marks on a femur shaft of a small young antelope.

BUT, they could have also been caused by: CROCODILES SLAMMING DOWN THEIR PREY
LIKE GODS OF DESTRUCTION

3.2 Ma
Au. Afarensis?
No actual tools found

Lomekwian Tools at 3.3 Ma?

• 2011, Kenya
• Large
• If truly tools, then oldest known
• But, no animal bones with cutmarks

Probable food items of Australopithecus:

Fruits
Seeds
Tubers
Some meat

AFTER ~ 1.9 MA THERE ARE NO MORE AUSTRALOPITHECUS OR KENYANTHROPUS

SPECIES
(me when the gummy starts hitting)

Paranthropus robustus
Robert Broom, others at on-going S. African sites Dated to 1.5 to 1.9mya

Paranthropus boisei
Louis Leakey, Mary Leakey, ongoing excavations
Sites include Olduvai Gorge, Koobi Fora (East Africa)
Dated to 1.0 to 2.3mya

Paranthropus aethiopicus
Alan Walker and colleagues
Sites include Omo Group and West of Lake Turkana
Dated to 2.5 to 2.3 mya (older than boisei & robustus)

The premolars look and function like molars

Sagittal crest in males and low, sloping frontal bone
Changes related to the temporalis muscle in Paranthropus species:
Large sagittal crest increases area for muscle attachment

Dietary Adaptations
• Stronger muscles generate larger forces.
• Anterior pillars support and absorb
stresses put on the area below the orbits

Large browridges resist bending stresses transferred upward from anterior pillars

Masseter muscle – between

zygomatic arch and lower jaw

A flat face means that muscle

attachment is closer to where the
food is to be crushed = greater bite force

• Paranthropus went
extinct around 1.4 Ma
• No living
descendants

Why?
• Too specialized?
• Climate change?
• Out-competed?

3/23

Louis and Mary Leakey

Paranthropus boisei (“Nutcracker Man”) very well preserved
Found in Olduvai Gorge

Leakeys looked for an alternative because...

Tools were probably used for meat processing, and Paranthropus was not a meat-eater
Tool making would seem to require a large brain, and the brain of Paranthropus wasn’t big
enough.

Olduvai Gorge 1960

Homo habilis
-found several of these specimen well preserved over the next decades

Every fossil found gets a unique id number

Homo habilis: Anatomy

(very small)
Cranial Capacity: 610cc
More prominent forehead
Brow ridges
Small face
Orthognathic face
Small dentition

Comparison of Australopithecus & Homo habilis

Australopithecus: Cranial capacity 450 cc (South Africa)
Homo Habilis: Cranial capacity 610 cc (Brow ridges Small face) (East Africa)

Postcranial Anatomy
-Small body size
(like Australopithecus)

Foot specialized for bipedalism

Retains relatively long arms

Hand bones indicate some climbing

Biggest differences between Homo habilis and Australopithecus are in

the skull

Homo rudolfensis
-In 1972 Richard Leakey found this skull,
KNM ER 1470

Homo rudolfensis
Cranial Capacity: 750cc
More Prominent Forehead
Small Brow Ridges
Large Face
Large Molars
Orthognathic Face

Two distinct species, or one, sexually dimorphic species?

Fossils that changed scientific perspective ... Part 1

(KNM-ER 1470)
-didn’t have teeth hard to directly compare it to remains
-Virtually complete skull of H. rudolfensis
-Different from H. habilis in shape
-2009 found mandible and some teeth

Part 2
Chelachew Seyoum and Kaye Reed
Ledi-Geraru, Ethiopia (ASU-led project)

-More similar to H.rudolfensis than to H. habilis

-Earliest fossil with Homo-like features
-2.8 Ma
-More evidence that H. habilis and H. rudolfensis are different species

Ecology/Diet
1. Australopithecus species had larger cheek teeth, thicker enamel and more massive jaws than
Miocene apes.
2. Paranthropus continued this trend with even larger cheek teeth and extremely massive jaws.
3. Homo reverses that trend with smaller cheek teeth and more delicate mandibles.
WHY???

Stone Tools!!!
o First evidence from ~ 2.5 Ma
o Made and used widely by 2.3 Ma in East Africa and ~2.0 Ma in South Africa
Oldowan Tools: (named after Olduvai Gorge, where they were found)
-fist-sized cobblestones, when
broken or chipped yielded a
usable core and several flakes

Were these tools used for hunting?

• Body size estimates for early Homo were around 52 kg (120 lbs) for males and 32 kg (70 lbs)
for females.
• Did these 70 to 120 pound hominins hunt by throwing these flakes?
o Probably not
• Were these types of flakes or cobbles attached to sticks?
o No evidence of this

What about power scavenging?

• Would these early Homo individuals be able to confront large predators AND drive them off?
• Probably not

Or, will they slink in after all the other scavengers are finished?

Evidence from animal fossils

• Cut marks only
- Animal deaths followed by Homo scavenging
• Cut marks beneath tooth marks
- Scavenging by Homo, followed by carnivore scavenging
• Cut marks above tooth marks
- Animals killed by carnivores followed by scavenging by Homo

So, what’s most likely?

• Early Homo probably wasn’t obtaining big game via hunting or power scavenging
o Small body size and lack of effective weapons
• Probably passive scavenging, including extracting marrow from long bones and brains from
skulls
• Also hunting of small prey

Late Pliocene Hominins

• Dietary morphology in Homo compared to Australopithecus
o Reduced mandible size
o Large incisors (unlike Paranthropus)
o Smaller molars and premolars

Possible food items of the Homo lineage

-Fruits
-Meat (more than Australopiths)
- Insects
Explains reduction in tooth size?

Homo erectus
• Known from 1.9 Ma - ~150,000y
• “Firsts”
o Hominins to leave the continent of Africa
o Fire, hearths, endurance running, etc.
o Modern human-like limb proportions
o Brains ~1000 cm 3

In 1891, along the banks of the Solo River...first hominins found outside Europe & Africa
-Eugene Dubois
-Java, Indonesia

Dubois found:
A skullcap, and a femur...
- clearly hominin – large cranial capacity
- clearly bipedal
Named it: Pithecanthropus erectus “upright ape man”
Ultimately, species changed to “Homo erectus”

Since Dubois, many specimens found on Java

Oldest: 1.6 Ma

AFRICAN HOMO ERECTUS

Homo erectus Koobi Fora, Keyna

-1.9 Ma

Homo Erectus Swartkrans, South Africa

Comparison of: African H. erectus vs. Asian H. erectus

African H: No Sagittal keel Asian H: Sagittal keel
Smaller brow-ridge Larger brow-ridge
-Smaller avg. brain size
(700-900cc range) -Larger avg. brain size(800-1000cc range)

Nariokotome Boy (Really complete skeleton)

African H. erectus
-Lake Turkana
-KNM-WT 15000
-8-9 yrs. old at death 5’ – 5’4” tall!
-Near modern body proportions
-Long legs relative to arms
-Narrow pelvis

Why is Homo erectus the first hominin to be found outside of Africa?

Acheulean Industry
-“Bifacial” - core is flaked along two distinct sides
- teardrop-shaped “handaxes

-Consistent proportions across time and space; similar proportions (Height:Width:Thickness),

regardless of size
• Pointed end cuts through meat, rounded end fits in palm

-Hand axes best work best for disarticulating bones and heavy cutting butchering

Controlled use of Fire?

-Stone tools found near baked earth
• Burned animal bones found with H. erectus fossils
-Experiments conducted to assess how soil and bone look after different kinds of fires
• Results suggest camp fires (rather than natural fires)

Selection Pressures in the Savanna that

Might Favor Larger Body Size and Obligate Bipedalism:
-More predators
• Greater distances to travel among resources

Homo erectus
-Dmanisi
-1.75 Ma
-600 – 775cc (H. habilis range)
• Challenges the notion that big brains propelled the exodus of early humans out of Africa

Dmanisi: Stone Tools

-Most similar to Oldowan tools from Olduvai...
-Simple chopping and scraping tools.

What came after Erectus?

“Archaic Homo sapiens”
• Group of hominins with
many overlapping features
• Not quite “modern”, but derived compared to H. erectus
• Historically separated into species based on geographic region, e.g.
– Africa: H. rhodesiensis
– Europe: H. heidelbergensis,
H. antecessor
– Asia: H. daliensis
a.k.a. “The muddle in the middle Pleistocene”

In this class, we will refer to these archaic H. sapiens as H. heidelbergensis

Different from H. erectus in that

 ● Larger brain
● More globular brain case – reduced postorbital constriction
● Generally no sagittal keel
● Decrease in cheek teeth size

H. heidelbergensis in Africa: Bodo, Ethiopia

• one of the earliest specimens (~600ka)
• Primitive features
– more prognathic face
– slight sagittal keel
– flat, receding forehead

Asian H. heidelbergensis
China & Indonesia
• >100 - 210 ka
• Mix of traits, but
different from
African H.
Heidelbergensis

H. erectus-like:
– thick cranial bones
– sagittal keel
– flat, receding frontal
bone
– low cranial vault

Derived
– short and small face
– rounded brain case
– reduced postorbital
constriction
– Large brain (~1200 cc)

Schöningen Spears

3 wooden throwing
spears, Germany
• ~400 ka
• Oldest completely
preserved hunting
weapons, along with
10 butchered horses
• Thus, first strong
evidence of large-
game hunting

Dentition
– Relatively small
molars
– large front teeth,
which are well
worn
Stone Tools
– Compared to H. erectus,
more cutting edge per
pound of flint
– Evidence of attachment
to shafts
– Carried flint from far away
– Also bone & ivory tools

100% of adults (male and female) >30y have healed fractures

Did Neandertals have language?

• We don’t know
• They probably could not make all the same sounds as us
• Language is symbolic – not tied to particular sounds